Gap formation and dynamics after long‐term steady state in an old‐growth Picea abies stand in Norway: Above‐ and belowground interactions

Stand dynamics and the gap initiation prior to gap formation are not well‐understood because of its long‐term nature and the scarcity of late‐successional stands. Reconstruction of such disturbance is normally based on historical records and dendroecological methods. We investigated gap initiation and formation at the fine‐scale stand level in the old‐growth reserve of Karlshaugen in Norway. Given its long‐term conservation history, and thorough mapping in permanent marked plots with spatially referenced trees, it provides an opportunity to present stand development before, during, and after gap formation. Late‐successional decline in biomass was recorded after more than 50 years of close to steady state. Gaps in the canopy were mainly created by large old trees that had been killed by spruce bark beetles. Snapping by wind was the main reason for treefall. Long‐term dominance of Norway spruce excluded downy birch and Scots pine from the stand. Comparisons of the forest floor soil properties between the gap and nongap area showed significantly higher concentrations of plant available Ca within the gap area. Plant root simulator (PRS?) probes showed significantly higher supply rates for Ca and Mg, but significantly lower K for the gap compared to the nongap area. Soil water from the gap area had significantly higher C:N ratios compared to the nongap area. Fine‐scale variation with increasing distance to logs indicated that CWD is important for leaking of DOC and Ca. Our long‐term study from Karlshaugen documents gap dynamics after more than 50 years of steady state and a multiscale disturbance regime in an old‐growth forest. The observed disturbance dynamic caused higher aboveground and belowground heterogeneity in plots, coarse woody debris, and nutrients. Our study of the nutrient levels of the forest floor suggest that natural gaps of old‐growth forest provide a long‐lasting biogeochemical feedback system particularly with respect to Ca and probably also N. Norway spruce trees near the gap edge responded with high plasticity to reduced competition, showing the importance of the edge zone as hot spots for establishing heterogeneity, but also the potential for carbon sequestration in old‐growth forest.     

Shifts in priming partly explain impacts of long‐term nitrogen input in different chemical forms on soil organic carbon storage

Input of labile organic carbon can enhance decomposition of extant soil organic carbon (SOC) through priming. We hypothesized that long‐term nitrogen (N) input in different chemical forms alters SOC pools by altering priming effects associated with N‐mediated changes in plants and soil microbes. The hypothesis was tested by integrating field experimental data of plants, soil microbes and two incubation experiments with soils that had experienced 10 years of N enrichment with three chemical forms (ammonium, nitrate and both ammonium and nitrate) in an alpine meadow on the Tibetan Plateau. Incubations with glucose–¹³C addition at three rates were used to quantify effects of exogenous organic carbon input on the priming of SOC. Incubations with microbial inocula extracted from soils that had experienced different long‐term N treatments were conducted to detect effects of N‐mediated changes in soil microbes on priming effects. We found strong evidence and a mechanistic explanation for alteration of SOC pools following 10 years of N enrichment with different chemical forms. We detected significant negative priming effects both in soils collected from ammonium‐addition plots and in sterilized soils inoculated with soil microbes extracted from ammonium‐addition plots. In contrast, significant positive priming effects were found both in soils collected from nitrate‐addition plots and in sterilized soils inoculated with soil microbes extracted from nitrate‐addition plots. Meanwhile, the abundance and richness of graminoids were higher and the abundance of soil microbes was lower in ammonium‐addition than in nitrate‐addition plots. Our findings provide evidence that shifts toward higher graminoid abundance and changes in soil microbial abundance mediated by N chemical forms are key drivers for priming effects and SOC pool changes, thereby linking human interference with the N cycle to climate change.     

Modelling environmental and survey influences on lek attendance using long‐term lek survey data

Targeted lek surveys, in which historically active lekking areas are surveyed over multiple years, are commonly used to monitor lek‐mating birds. Lek surveys are usually used in habitats and species in which individuals are not easily detected when they are not lekking. Unfortunately, lek surveys suffer from a number of shortcomings, most notably the failure to observe lekking individuals when they are near but not actively attending a lek. Like other sources of imperfect detection, inconsistency in lek attendance may bias estimates of annual population sizes or the proportion of lekking arenas active in a given year. In turn, this may result in unreliable estimates of trends, distributions or other parameters of interest. An improved understanding of the factors affecting lek attendance is crucial to improving the efficacy of many lek monitoring programmes and the utility of lek surveys in estimating population and distributional dynamics. We assess variation in lek attendance in the Lesser Prairie Chicken Tympanuchus pallidicinctus using a subset of data from a long‐term monitoring programme in eastern New Mexico, USA. Focusing on lek‐years in which at least one male was observed, we used generalized linear models to assess the influence of lek size and survey and environmental characteristics on the probability that at least one male was observed in attendance. Our best‐supported model provided evidence for influences of each of these sources of variation. Models suggested that surveyors may improve the likelihood of detecting lek attendance by making small changes to survey duration, targeted survey dates and by considering environmental factors such as weather, habitat and human disturbance. In addition to improving detection, consideration of lek attendance patterns and associated behaviours may improve estimation for more effective conservation in a variety of lekking bird species.     

Plant diversity positively affects short‐term soil carbon storage in experimental grasslands

Increasing atmospheric CO₂ concentration and related climate change have stimulated much interest in the potential of soils to sequester carbon. In ‘The Jena Experiment’, a managed grassland experiment on a former agricultural field, we investigated the link between plant diversity and soil carbon storage. The biodiversity gradient ranged from one to 60 species belonging to four functional groups. Stratified soil samples were taken to 30 cm depth from 86 plots in 2002, 2004 and 2006, and organic carbon contents were determined. Soil organic carbon stocks in 0–30 cm decreased from 7.3 kg C m^?2 in 2002 to 6.9 kg C m^?2 in 2004, but had recovered to 7.8 kg C m^?2 by 2006. During the first 2 years, carbon storage was limited to the top 5 cm of soil while below 10 cm depth, carbon was lost probably as short‐term effect of the land use change. After 4 years, carbon stocks significantly increased within the top 20 cm. More importantly, carbon storage significantly increased with sown species richness (log‐transformed) in all depth segments and even carbon losses were significantly smaller with higher species richness. Although increasing species diversity increased root biomass production, statistical analyses revealed that species diversity per se was more important than biomass production for changes in soil carbon. Below 20 cm depth, the presence of one functional group, tall herbs, significantly reduced carbon losses in the beginning of the experiment. Our analysis indicates that plant species richness and certain plant functional traits accelerate the build‐up of new carbon pools within 4 years. Additionally, higher plant diversity mitigated soil carbon losses in deeper horizons. This suggests that higher biodiversity might lead to higher soil carbon sequestration in the long‐term and therefore the conservation of biodiversity might play a role in greenhouse gas mitigation.     

Short‐ and long‐term costs of reproduction in a migratory songbird

Costs of reproduction represent a common life‐history trade‐off. Critical to understanding these costs in migratory species is the ability to track individuals across successive stages of the annual cycle. We assessed the effects of total number of offspring fledged and date of breeding completion on pre‐migratory body condition, the schedule of moult and annual survival in a migratory songbird, the Savannah Sparrow Passerculus sandwichensis. Between 2008 and 2010, moult was delayed for individuals that finished breeding later in the breeding period and resulted in reduced lean tissue mass during the pre‐migratory period, suggesting an indirect trade‐off between the timing of breeding completion and condition just prior to migration. Lean tissue mass decreased as the number of offspring fledged increased in 2009, a particularly cool and wet year, illustrating a direct trade‐off between reproductive effort and condition just prior to migration in years when weather is poor. However, using a 17‐year dataset from the same population, we found that parents that fledged young late in the breeding period had the highest survival and that number of offspring fledged did not affect survival, suggesting that individuals do not experience long‐term trade‐offs between reproduction and survival. Taken together, our results suggest that adult Savannah Sparrows pay short‐term costs of reproduction, but that longer‐term costs are mitigated by individual quality, perhaps through individual variation in resource acquisition.     

The contrasting effects of short‐ and long‐term habitat drainage on the population dynamics of freshwater turtles in a human‐dominated landscape

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Influence of long‐term land application of Class B biosolids on soil bacterial diversity

Aim: To evaluate the effect of long‐term annual land applications of Class B biosolids on soil bacterial diversity at University of Arizona Marana Agricultural Field Center, Tucson, Arizona. Methods and Results: Following the final of 20 consecutive years of application of Class B biosolids in March 2005, followed by cotton growth from April to November 2005 surface soil samples (0–30 cm) were collected from control (unamended) and biosolid‐amended plots. Total bacterial community DNA was extracted, amplified using 16S rRNA primers, cloned, and sequenced. All 16S rRNA sequences were identified by 16S rRNA sequence analysis and comparison to known sequences in GenBank (NCBI Blast N and Ribosomal Database Project II, RDP). Results showed that the number of known genera (identifiable > 96%) increased in the high rate biosolid plots compared to control plots. Biosolids‐amended soils had a broad phylogenetic diversity comprising more than four major phyla: Proteobacteria (32%), Acidobacteria (21%), Actinobacteria (16%), Firmicutes (7%), and Bacteroidetes (6%) which were typical to bacterial diversity found in the unamended arid southwestern soils. Conclusion: Bacterial diversity was either enhanced or was not negatively impacted following 20 years of land application of Class B biosolids. Significance and Impact of the Study: This study illustrates that long‐term land application of biosolids to arid southwestern desert soils has no deleterious effect on soil microbial diversity.     

Short‐term and long‐term effects of acute kidney injury in chronic kidney disease patients: A longitudinal analysis

We use data from an ongoing cohort study of chronic kidney patients at Salford Royal NHS Foundation Trust, Greater Manchester, United Kingdom, to investigate the influence of acute kidney injury (AKI) on the subsequent rate of change of kidney function amongst patients already diagnosed with chronic kidney disease (CKD). We use a linear mixed effects modelling framework to enable estimation of both acute and chronic effects of AKI events on kidney function. We model the fixed effects by a piece‐wise linear function with three change‐points to capture the acute changes in kidney function that characterise an AKI event, and the random effects by the sum of three components: a random intercept, a stationary stochastic process with Matérn correlation structure, and measurement error. We consider both multivariate Normal and multivariate t versions of the random effects. For either specification, we estimate model parameters by maximum likelihood and evaluate the plug‐in predictive distributions of the random effects given the data. We find that following an AKI event the average long‐term rate of decline in kidney function is almost doubled, regardless of the severity of the event. We also identify and present examples of individual patients whose kidney function trajectories diverge substantially from the population‐average.     

Restorative mowing on an abandoned semi‐natural meadow: short‐term and predicted long‐term effects

Abstract. When management, in the form of cattle grazing and mowing, ceases the abundance of competitively superior plant species tends to increase in abandoned semi‐natural meadows. Litter accumulation elevates the soil nutrient levels and hinders seedling recruitment. We surveyed changes in plant cover and species composition of a formerly grazed meadow in permanent plots for six years. Some plots were unmown, while others were mown and raked annually in August. The cover of grasses decreased and herb cover remained unchanged regardless of the treatment. Mowing and raking significantly reduced litter accumulation and increased the number of ground layer species. The expected long‐term effects of abandonment and restorative mowing were studied by calculating the transition probabilities for unmown and mown plots and simulating the course of succession as projected by the transition matrices. During a simulation period of 30 yr, abandonment led to (1) a decrease in the cover of small herbs, (2) a slight increase in the cover of tall herbs and (3) a slight decrease in the cover of grasses. In contrast, the cover of small herbs on the mown plots remained unchanged or slightly increased during the course of simulation. These results suggest that mowing late in the season is primarily a management tool for the maintenance of the existing species diversity and composition. However, it may not be an effective restorative tool to induce overall changes in the resident vegetation of abandoned grass‐dominated meadows. Grazing or mowing early in the season may be more effective in this respect. Consequently, mowing early or, alternatively, late in the season may provide management strategies for the maintenance and restoration of species diversity, respectively.     

A survey of long‐term terrestrial ecology studies in Australia

Long‐term ecological studies (LTES) are critical for understanding and managing landscapes. To identify important research gaps, facilitate collaborations and communicate results, several countries have established long‐term ecological research networks. A few initiatives to create such a network in Australia have been undertaken, but relatively few published data exist on the current state of LTES in Australia. In this paper, we present the results of an online survey of terrestrial LTES projects sent to academic, government and non‐governmental organization‐based researchers across Australia. We asked questions pertaining to the focus, scope, support and outcomes of LTES spanning 7 years or longer. Based on the information reported from 85 Australian LTES, we: (i) identify the biomes, processes and species that are under‐represented in the current body of research; (ii) discuss important contributing factors to the successful development and survival of these projects; and (iii) make recommendations to help increase the productivity and influence of LTES across research, management and policy sectors.     

Shift in community structure in an early‐successional Mediterranean shrubland driven by long‐term experimental warming and drought and natural extreme droughts

Global warming and recurring drought are expected to accelerate water limitation for plant communities in semiarid Mediterranean ecosystems and produce directional shifts in structure and composition that are not easily detected, and supporting evidence is scarce. We conducted a long‐term (17 years) nocturnal‐warming (+0.6°C) and drought (?40% rainfall) experiments in an early‐successional Mediterranean shrubland to study the changes in community structure and composition, contrasting functional groups and dominant species, and the superimposed effects of natural extreme drought. Species richness decreased in both the warming and drought treatments. Responses to the moderate warming were associated with decreases in herb abundance, and responses to the drought were associated with decreases in both herb and shrub abundances. The drought also significantly decreased community diversity and evenness. Changes in abundance differed between herbs (decreases) and shrubs (increases or no changes). Both warming and drought, especially drought, increased the relative species richness and abundance of shrubs, favoring the establishment of shrubs. Both warming and drought produced significant shifts in plant community composition. Experimental warming shifted the community composition from Erica multiflora toward Rosmarinus officinalis, and drought consistently shifted the composition toward Globularia alypum. The responses in biodiversity (e.g., community biodiversity, changes of functional groups and compositional shifts) were also strongly correlated with atmospheric drought (SPEI) in winter–spring and/or summer, indicating sensitivity to water limitation in this early‐successional Mediterranean ecosystem, especially to natural extreme droughts. Our results suggest that the shifts in species assembles and community diversity and composition are accelerated by the long‐term nocturnal‐warming and drought, combined with natural severe droughts, and that the magnitude of the impacts of climate change is also correlated with the successional status of ecosystem. The results thus highlight the necessity for assessing the impacts on ecosystemic functioning and services and developing effective measures for conserving biodiversity.     

Dampening prey cycle overrides the impact of climate change on predator population dynamics: a long‐term demographic study on tawny owls

Predicting the dynamics of animal populations with different life histories requires careful understanding of demographic responses to multifaceted aspects of global changes, such as climate and trophic interactions. Continent‐scale dampening of vole population cycles, keystone herbivores in many ecosystems, has been recently documented across Europe. However, its impact on guilds of vole‐eating predators remains unknown. To quantify this impact, we used a 27‐year study of an avian predator (tawny owl) and its main prey (field vole) collected in Kielder Forest (UK) where vole dynamics shifted from a high‐ to a low‐amplitude fluctuation regime in the mid‐1990s. We measured the functional responses of four demographic rates to changes in prey dynamics and winter climate, characterized by wintertime North Atlantic Oscillation (wNAO). First‐year and adult survival were positively affected by vole density in autumn but relatively insensitive to wNAO. The probability of breeding and number of fledglings were higher in years with high spring vole densities and negative wNAO (i.e. colder and drier winters). These functional responses were incorporated into a stochastic population model. The size of the predator population was projected under scenarios combining prey dynamics and winter climate to test whether climate buffers or alternatively magnifies the impact of changes in prey dynamics. We found the observed dampening vole cycles, characterized by low spring densities, drastically reduced the breeding probability of predators. Our results illustrate that (i) change in trophic interactions can override direct climate change effect; and (ii) the demographic resilience entailed by longevity and the occurrence of a floater stage may be insufficient to buffer hypothesized environmental changes. Ultimately, dampened prey cycles would drive our owl local population towards extinction, with winter climate regimes only altering persistence time. These results suggest that other vole‐eating predators are likely to be threatened by dampening vole cycles throughout Europe.     

Topological effects of network structure on long‐term social network dynamics in a wild mammal

Social structure influences ecological processes such as dispersal and invasion, and affects survival and reproductive success. Recent studies have used static snapshots of social networks, thus neglecting their temporal dynamics, and focused primarily on a limited number of variables that might be affecting social structure. Here, instead we modelled effects of multiple predictors of social network dynamics in the spotted hyena, using observational data collected during 20 years of continuous field research in Kenya. We tested the hypothesis that the current state of the social network affects its long‐term dynamics. We employed stochastic agent‐based models that allowed us to estimate the contribution of multiple factors to network changes. After controlling for environmental and individual effects, we found that network density and individual centrality affected network dynamics, but that social bond transitivity consistently had the strongest effects. Our results emphasise the significance of structural properties of networks in shaping social dynamics.     

Functional traits determine tree growth and ecosystem productivity of a tropical montane forest: Insights from a long‐term nutrient manipulation experiment

Trait‐response effects are critical to forecast community structure and biomass production in highly diverse tropical forests. Ecological theory and few observation studies indicate that trees with acquisitive functional traits would respond more strongly to higher resource availability than those with conservative traits. We assessed how long‐term tree growth in experimental nutrient addition plots (N, P, and N + P) varied as a function of morphological traits, tree size, and species identity. We also evaluated how trait‐based responses affected stand scale biomass production considering the community structure. We found that tree growth depended on interactions between functional traits and the type or combination of nutrients added. Common species with acquisitive functional traits responded more strongly to nutrient addition, mainly to N + P. Phosphorous enhanced the growth rates of species with acquisitive and conservative traits, had mostly positive effects on common species and neutral or negative effects in rare species. Moreover, trees receiving N + P grew faster irrespective of their initial size relative to trees in control or to trees in other treatment plots. Finally, species responses were highly idiosyncratic suggesting that community processes including competition and niche dimensionality may be altered under increased resource availability. We found no statistically significant effects of nutrient additions on aboveground biomass productivity because acquisitive species had a limited potential to increase their biomass, possibly due to their generally lower wood density. In contrast, P addition increased the growth rates of species characterized by more conservative resource strategies (with higher wood density) that were poorly represented in the plant community. We provide the first long‐term experimental evidence that trait‐based responses, community structure, and community processes modulate the effects of increased nutrient availability on biomass productivity in a tropical forest.     

A method for the automated long‐term monitoring of three‐spined stickleback Gasterosteus aculeatus shoal dynamics

This paper describes and evaluates a flexible, non‐invasive tagging system for the automated identification and long‐term monitoring of individual three‐spined sticklebacks Gasterosteus aculeatus. The system is based on barcoded tags, which can be reliably and robustly detected and decoded to provide information on an individual's identity and location. Because large numbers of fish can be individually tagged, it can be used to monitor individual‐ and group‐level dynamics within fish shoals.     

Long‐term colonization ecology of forest‐dwelling species in a fragmented rural landscape – dispersal versus establishment

Species colonization in a new habitat patch is an efficiency indicator of biodiversity conservation. Colonization is a two‐step process of dispersal and establishment, characterized by the compatibility of plant traits with landscape structure and habitat conditions. Therefore, ecological trait profiling of specialist species is initially required to estimate the relative importance of colonization filters. Old planted parks best satisfy the criteria of a newly created and structurally matured habitat for forest‐dwelling plant species. We sampled species in 230 ancient deciduous forests (source habitat), 74 closed‐canopy manor parks (target habitats), 151 linear wooded habitats (landscape corridors), and 97 open habitats (isolating matrix) in Estonia. We defined two species groups of interest: forest (107 species) and corridor specialists (53 species). An extra group of open habitat specialists was extracted for trait scaling. Differing from expectations, forest specialists have high plasticity in reproduction mechanisms: smaller seeds, larger dispersules, complementary selfing ability, and diversity of dispersal vectors. Forest specialists are shorter, less nutrient‐demanding and mycorrhizal‐dependent, stress‐tolerant disturbance‐sensitive competitors, while corridor specialists are large‐seeded disturbance‐tolerant competitors. About 40% of species from local species pools have immigrated into parks. The historic forest area, establishment‐related traits, and stand quality enhance the colonization of forest specialists. The openness of landscape and mowing in the park facilitate corridor specialists. Species traits in parks vary between a forest and corridor specialist, except for earlier flowering and larger propagules. Forest species are not dispersal limited, but they continue to be limited by habitat properties even in the long term. Therefore, the shady parts of historic parks should be appreciated as important forest biodiversity‐enhancing landscape structures. The habitat quality of secondary stands can be improved by nurturing a heterogeneous shrub and tree layer, and modest herb layer management.